CSIRO PUBLISHING

Wildlife Research, 2022, 49, 295–302

https://doi.org/10.1071/WR21057

Turning ghosts into dragons: improving camera monitoring

outcomes for a cryptic low-density Komodo dragon
population in eastern Indonesia

Deni Purwandana A, Achmad Ariefiandy A, Muhammad Azmi A,

Sanggar A. Nasu A, Sahudin B, Andreas A. Dos B and Tim S. Jessop A,C,D

A
Komodo Survival Program, Jl. Karang Sari Blok G, No. 10, Denpasar 80117, Bali, Indonesia.
B
Balai Besar Konservasi Sumber Daya Alam Nusa Tenggara Timur, Jl. SK Lerik kelapa lima, Kupang 85228,
NTT, Indonesia.
C
Centre for Integrative Ecology, School of Life and Environmental Sciences, Deakin University, 75 Pigdons
Road, Waurn Ponds, Vic. 3220, Australia.
D
Corresponding author. Email: t.jessop@deakin.edu.au

Abstract
Context. Detection probability is a key attribute influencing population-level wildlife estimates necessary for
conservation inference. Increasingly, camera traps are used to monitor threatened reptile populations and communities.
Komodo dragon (Varanus komodoensis) populations have been previously monitored using camera traps; however,
considerations for improving detection probability estimates for very low-density populations have not been well
investigated.
Aims. Here we compare the effects of baited versus non-baited camera monitoring protocols to influence Komodo
dragon detection and occupancy estimates alongside monitoring survey design and cost considerations for ongoing
population monitoring within the Wae Wuul Nature Reserve on Flores Island, Indonesia.
Methods. Twenty-six camera monitoring stations (CMS) were deployed throughout the study area with a minimum of
400 m among CMS to achieve independent sampling units. Each CMS was randomly assigned as a baited or non-baited
camera monitoring station and deployed for 6 or 30 daily sampling events.
Key results. Baited camera monitoring produced higher site occupancy estimates with reduced variance. Komodo
dragon detection probability estimates were 0.15
0.092–0.22 (95% CI), 0.01
0.001–0.03, and 0.03
0.01–0.04 for
baited (6 daily survey sampling events), unbaited (6 daily survey sampling events) and long-unbaited (30 daily survey
sampling events) sampling durations respectively. Additionally, the provision of baited lures at cameras had additional
benefits for Komodo detection, survey design and sampling effort costs.
Conclusions. Our study indicated that baited cameras provide the most effective monitoring method to survey low-
density Komodo dragon populations in protected areas on Flores.
Implications. We believe our monitoring approach now lends itself to evaluating population responses to ecological
and anthropogenic factors, hence informing conservation efforts in this nature reserve.

Keywords. population monitoring, effective sampling, protected areas, apex predator, reptiles, Varanus komodoensis.

Received 25 March 2021, accepted 13 July 2021, published online 16 December 2021

Introduction conservation actions on their populations (Karanth et al. 2004;

Large terrestrial predators are most often at risk from human
actions and increasingly require effective conservation actions
to ensure population persistence (Gittleman and Harvey 1982;
Prowse et al. 2015; Penjor et al. 2019). The key requirement to
establish effective conservation actions for apex predators is to
accurately monitor population trends and status (Karanth et al.
2011). However, because apex predators are often rare or averse
to capture or detect, non-invasive monitoring methods are rou-
tinely used to evaluate the effects of threatening processes or
O’Connell et al. 2010). Similarly, the increasing use of hierar-
chical models such as site occupancy and n-mixture models,
which account for imperfect detection, are now among the most
common techniques used to provide population-level inference
for apex predators (MacKenzie et al. 2002, 2006; Royle 2004;
Kéry et al. 2005). Indeed, these methods are often well suited for
threatened predator population studies (du Preez et al. 2014; Tan
et al. 2017; Penjor et al. 2019; Searle et al. 2020), because
threatened predators often persist at low densities where

Journal compilation Ó CSIRO 2022 Open Access CC BY www.publish.csiro.au/journals/wr

296 Wildlife Research
individual-based recapture or resighting probabilities can be too
low to allow for the alternate population estimates using mark–
recapture type models (Williams et al. 2002; Kéry and Schmidt
2008; Couturier et al. 2013; du Preez et al. 2014; Tan et al. 2017;
Searle et al. 2020).
Non-invasive monitoring techniques such as camera trapping
are now increasingly used for monitoring terrestrial reptiles, a
taxon with over 11 000 primarily predatory species (Ariefiandy
et al. 2013; Jessop et al. 2013; Welbourne et al. 2015; Adams
et al. 2017; Moore et al. 2020). Nevertheless, the use of cameras,
as measured by the capacity to achieve adequate detection for
robust population-level estimates, remains variable within and
among reptile species because of the effects of body size, species
habits and environmental factors (Ariefiandy et al. 2013;
Welbourne et al. 2015; Richardson et al. 2017; Einoder et al.
2018). In the case of large reptiles, lower population densities,
greater daily movement capacity, the effects of seasonal climatic
variation, and smaller skin surface to ambient air temperature
differences can all influence camera-based population monitoring
effectiveness (Ariefiandy et al. 2013; Jessop et al. 2013;
Welbourne 2013; Richardson et al. 2017; Hu et al. 2019).
Furthermore, human activities can often disproportionately
threaten large-bodied reptiles, causing their populations to be at
much lower densities than normal and thus more difficult to
monitor (Todd et al. 2010; Tingley et al. 2019). Thus, addressing
these factors by modifying camera sampling designs to increase
detection probability is a key consideration to monitor threatened
reptile populations effectively. Under such circumstances, there
may be compelling reasons to improve camera-based detection
using baits or lures (i.e. attractants) to increase detection proba-
bility (O’Connell et al. 2010; Long et al. 2012; Read et al. 2015).
Multiple studies have reported that the use of baits or lures as
attractants can vastly improve predators’ detection sensitivity
(du Preez et al. 2014; Austin et al. 2017; Comer et al. 2018). This
result is especially important in predator populations where
individuals can be cryptic or persist as low-density populations.
Hence, attractants or baits may be essential to increase detection
to prevent poor quality estimates of population-level parameters
(Thompson 2013). For this reason, baited camera traps deployed
during appropriate weather conditions can be advocated to
optimise large-reptile detection probability (Jessop et al.
2013). Although, it is important to note that these benefits
may need to consider how baits can affect a species’ movement
behaviour and create potential biases in any arising population-
level estimates (Stewart et al. 2019).
The Komodo dragon (Varanus komodoensis) is the largest
lizard and has an important ecological role as an apex predator
(Jessop et al. 2006, 2019, 2020). The current distribution of
Komodo dragons is restricted to five islands located in Komodo
National Park and several fragmented populations on Flores
Island (Jessop et al. 2007, 2018; Purwandana et al. 2014a;
Ariefiandy et al. 2015; Jones et al. 2020). Populations on Flores
Island have decreased because of anthropogenic activities and
are now increasingly reliant on a small number of reserve areas
to ensure their persistence (Ariefiandy et al. 2015, 2020; Jones
et al. 2020).
Komodo dragon populations on Flores persist at much lower
population densities (,1 dragon km2) than those observed in
Komodo National Park (,10 dragons km2; Laver et al. 2012;
D. Purwandana et al.
Purwandana et al. 2014a; Ariefiandy et al. 2015, 2020). Multi-
ple field methods have been used to estimate population trends
of Komodo dragons (Ariefiandy et al. 2013, 2014). However,
these can vary considerably in their monitoring effectiveness
(Jessop et al. 2007; Ariefiandy et al. 2013, 2014; Purwandana
et al. 2014a, 2015). On Flores, low densities and trap-wary
behaviour of Komodo dragons favour wildlife cameras over
direct trapping methods as a more effective population monitor-
ing methodology (Ariefiandy et al. 2015). However, optimising
camera monitoring design is still necessary to allow conserva-
tion managers to improve the data used to evaluate these most
vulnerable populations (Jones et al. 2020). Here, we compare the
effect of baited and unbaited camera sampling on the estimates
of Komodo dragon detection probability and site occupancy,
alongside other measures of monitoring efficacy and project
running costs within the Wae Wuul Nature Reserve of Flores.
Finally, we discuss the implications of our results for managing
Komodo dragons within this protected area and, more broadly,
for other populations distributed on the island of Flores.
Materials and methods
Study area
The Wae Wuul Nature Reserve comprises a protected area of
14.84 km2 located on the western coast of Flores in eastern
Indonesia (Fig. 1a, b). The reserve was established in 1985,
aiming to increase protection of Komodo dragons beyond
Komodo National Park. The climate is highly seasonal, domi-
nated by a long dry season from March to November and a short
wet season. Annual rainfall is less than 2000 mm (Monk et al.
1997). The study area comprises a hilly coastal landscape covered
in multiple distinct vegetation communities. The two most com-
mon vegetation communities are savanna grassland (common
species include Eulalia leschenaultiana and Setaria adhaerens)
and savanna woodland (common species include Borassus
flabellifer and Zizyphus horsfeldi) that cover ,80% of the study
area (Auffenberg 1981). In valley floors holding permanent or
ephemeral watercourses, drier vegetation communities are
replaced by open deciduous monsoon forest (,20% of the study
area; dominant species include Tamarindus indica, Schleichera
oleosa and Cassia javanica) or bamboo forest. These land cover
types are representative of those found across the lowland coastal
areas of major islands in this region of eastern Indonesia,
including the adjacent Komodo National Park (Auffenberg 1981).
Study design
Twenty-six camera monitoring stations (CMS) were deployed
within the Wae Wuul Nature Reserve. These CMS were placed
within all key vegetation communities, including deciduous
monsoon forest and savanna woodland. A minimum of 400 m
separated all sites to improve data independence obtained from
cameras (Ariefiandy et al. 2013, 2014). This 400-m distance
between monitoring sites was based on the radius of the mean
home-range area for Komodo dragons (Jessop et al. 2018;
Purwandana et al. 2021). At the commencement of the study,
each site was randomly assigned as a baited (n ¼ 13) or non-
baited (n ¼ 13) camera monitoring station to ensure equal
replicates within each camera method treatment. After the initial
sampling period at each station, the assigned camera method
Improving camera monitoring of Komodo dragons Wildlife Research 297

(a) (b)

(c)

Fig. 1. The study evaluated the effect of bait attractant on Komodo dragon detection probability and site occupancy estimates by using
camera monitoring stations deployed across (a) the Wae Wuul Nature Reserve located (b) on the western coast of Flores in eastern
Indonesia. (c) An image of a Komodo dragon inspecting the meat attractant contained within a metal bait box.

was reversed to the alternate method to compare estimates of
Komodo dragon detection probability and occupancy obtained
for each method at each site. The study was conducted during
June and July 2017 in the mid-dry season when environmental
temperatures permit Komodo dragons to exhibit normal daily
diurnal activity patterns and, hence, pending abundance, the
potential for good detection probability (Harlow et al. 2010a,
2010b; Jessop et al. 2013).
Camera monitoring design
At each CMS, a single outward facing Bushnell camera (Model
Trophy Cam HD 119678) was attached to a tree (40 cm above
the ground) as described elsewhere (Ariefiandy et al. 2013,
2014). Cameras were programmed to take three photos and a
1-min video each time an animal triggered the device. At
installation, all cameras were tested to confirm normal func-
tioning. For CMS allocated to the bait treatment, we used two
scent lures that comprised a small aluminium box (25  15  15
cm; L  W  H) and a suspended plastic bag, each containing
goat meat that was placed 4 and 2 m in front or above the camera
respectively. Baited and unbaited CMS were deployed for 6 and
30 days of monitoring respectively. The uneven durations
between treatments reflected our belief that baited cameras
would require considerably less sampling effort to produce
higher detection probabilities than those obtained from unbaited
cameras. As Komodo dragons have been observed to investigate
baits at traps for several minutes before entering traps or moving
elsewhere, we also used a 30-min camera delay to prevent
repeated photography of the same individual lizard (Ariefiandy
et al. 2014). In addition, a 3-day non-monitoring period was used
immediately after the transition from baited to unbaited sites.
This waiting period was implemented to remove a potential
carry-over bait effect that could have attracted Komodo dragons
and inflated detection probability at sites then monitored with
unbaited cameras. This research abided by the journal’s guide-
lines on ethical standards.
Estimating detection and site occupancy estimates
We modelled the detectability and occupancy of Komodo dra-
gons by using a single-season occupancy model, using the
298 Wildlife Research D. Purwandana et al.

Table 1. Model selection results testing effects of baited and non-baited cameras for influencing detection probability (p) and site occupancy (C) of
Komodo dragons within the Wae Wuul Nature Reserve in West Flores
K, the number of estimated parameters; logLik, logliklihood; AIC, Akaike information criterion; DAIC, the difference in value between AIC of this model and
the most parsimonious model; and AIC weights (wi), a measure of relative model support

Model K logLik AIC DAIC wi

C(.) p(bait vs no-bait)) 3 365.43 371.43 0.00 0.71
C(bait vs no-bait)) p(bait vs no-bait)) 4 365.21 373.21 1.78 0.29
C(.) p(bait vs no-bait))* daily survey variation 62 302.17 426.17 54.74 0.00
C (.) p (.) 2 424.44 428.44 57.01 0.00

software Presence (Hines 2006). Site occupancy models use
patterns of detection and non-detection over multiple surveys
(sampling occasions) of a sampling unit (CMS) to estimate
detection probabilities (p) and, thus, produce unbiased estimates
of occupancy (c) (MacKenzie et al. 2002). We modelled the
effect of baits on both the detection probability (p) and c relative
to those cameras without baits (i.e. p. c). We partitioned the
unbaited CMS detection probability data into two datasets,
given the sampling duration differences between baited and
unbaited CMS. One dataset comprised the first six, and the other
the full 30 daily sampling events. Models were ranked using
AIC, and we considered the effect of bait provision at CMS to be
influential if the model AIC was .2 units below that estimated
for the null model (Burnham and Anderson 2004).
Detection probability curves, probability of site absences
and survey design costs
To assess the expected reduction in sampling effort provided by
using baits at CMS, we produced detectability curves for CMS
with and without baits. Detectability curves represent the
cumulative probability (i.e. rate of increase) that Komodo dra-
gons will be detected after a given number of sampling occa-
sions in a site where the species is present (Wintle et al. 2005).
Cumulative detection probability curves were estimated as
pk ¼ 1 – (1 – p)k, where p is the species’ per-survey detection
probability within a given treatment and k is the given number of
sampling occasions (MacKenzie and Royle 2005).
Next, we estimated the minimum number of sequential
sampling occasions, with no detection required to be 95%
certain (i.e. a ¼ 0.05) that Komodo dragons were absent from
a surveyed site by using baited and unbaited cameras (Wintle
et al. 2012; Ferreras et al. 2018). The probability (with a ¼ 0.05)
of not detecting Komodo dragons after N sampling occasions at
a given site is estimated by the formula

a

C 
log  log 1C
N> 1a
logð1  pÞ
X
C ðmÞ ¼ ðCd þ Cr þ Cb  Sd þ Cbb  Sd þ Ccb  Sd Þ
where C(m) ¼ method specific survey cost, Cd ¼ cost of camera
deployment, Cr ¼ cost of camera retrieval, Cb ¼ cost of bait
(US$0.26 per camera per survey day (Sd)), Cbb ¼ cost of bait
boxes (US$10.00 per camera), Ccb ¼ cost of camera batteries
(US$0.20 per camera per survey day).
Results
The most parsimonious occupancy model (C (.), p (bait vs no-
bait), model weight ¼ 0.71) indicated that the effect of baits
placed at CMS vastly improved Komodo dragon detection
probability compared with the null model (DAIC ¼ 57.01; model
weight ¼ 0.00; Table 1). Detection probability estimates for
baited cameras (6 daily sampling events) were 15 and 5.5 times
higher than those estimated for unbaited (6 daily sampling events)
and long-unbaited (30 daily sampling events) camera sampling
durations (Fig. 2a). Similarly, baited cameras produced 2.3 and
1.3 higher Komodo dragon site occupancy estimates at the
equivalent and long-unbaited camera sampling durations
(Fig. 2b). A goodness-of-fit test on the most parameter-rich model
demonstrated that our data were not over-dispersed (i.e. ĉ . 1).
Effects of bait attractants on monitoring considerations
Baited cameras improved sampling efficacy and reduced mon-
itoring costs compared with sampling using unbaited cameras.
First, it was evident that based on cumulative detection proba-
bilities, baited cameras, if deployed sufficiently long enough,
could achieve perfect detection at sites with Komodo dragons,
with much less survey effort than with unbaited cameras (Fig. 3).
Compared with unbaited cameras, baited cameras reduced the
sampling effort duration from 184 to 21 days to be certain (with
a of ,0.05) that Komodo dragons were absent from a site.
Finally, because of the much-improved detection probability
achieved with baited cameras, it reduced the overall study costs
from US$580.20 to US$547.60, to obtain similar camera-based
detection levels within the study area.

Here, values of p and c are specific to baited and unbaited Discussion

CMS site occupancy estimates derived from the 6-day sampling
period.
Finally, we compared the costs of sampling for baited and
unbaited camera trapping methodology to achieve a similar
monitoring outcome (i.e. a ¼ 0.05) by calculating protocol-
specific costs of each technique, beyond common costs associ-
ated with camera purchases, as such we estimated
The choice of an appropriate sampling method for monitoring
threatened predator populations depends on interactions among
the program objectives, scale and resources and a species’s
detection probability (Kéry and Schmidt 2008). We demon-
strated that using baited-camera compared with unbaited-
camera monitoring greatly improved estimates of Komodo
dragon detection probability and site occupancy in the Wae
Improving camera monitoring of Komodo dragons Wildlife Research 299

0.25
(a) Wuul Nature Reserve on Flores Island. Indeed, several clear

Komodo dragon detection probability

advantages were evident from using baited cameras, including a
0.20 reduced sampling effort and, ultimately, a more cost-effective
monitoring design.
0.15
Obtaining a high detection probability is a key requirement to
improve site-occupancy estimates for large predators that persist
at low density (MacKenzie et al. 2006). Such sampling designs
0.10 should aim to achieve a detection probability exceeding 0.15, so
as to allow for better occupancy estimates for predators
0.05
(O’Connell et al. 2010; Otto and Roloff 2011). With camera-
based monitoring, there are several ways to increase species
detection probability, including increasing the number of cam-
0.00 eras deployed for longer survey periods or by also placing
d (6) aited (6) ited (
30) cameras in areas that increase detection opportunities of the
Baite Unb Unba
focal species (e.g. along game trails; O’Connell et al. 2010;
Camera method Geyle et al. 2020; Wysong et al. 2020). However, the use of
1.0
attractants such as baits or lures is another common means to
(b) improve camera-based detection probability, but their use should
Komodo dragon site occupancy

be assessed to ensure improved efficacy (Read et al. 2015).

0.8 It was evident that bait attractants at camera monitoring
stations greatly improved Komodo dragon detection probability
0.6
by 3.5–5 times over similar or extended durations of unbaited
camera monitoring. This finding is consistent with those of other
studies that indicate similar benefits of using baits or lures at
0.4 camera monitoring stations (du Preez et al. 2014; Austin et al.
2017; Tarugara et al. 2019). Importantly, these gains in detec-
0.2 tion probability alongside higher and more robust estimates of
site occupancy offset the increased daily sampling costs owing
to the purchase of goats as the bait source (Thorn et al. 2011).
0.0 Another key benefit of baited cameras was the considerable
d (6) aited (6) ited (
30) reduction (i.e. 5-fold) in the survey effort needed to achieve
Baite Unb Unba
adequate Komodo dragon detection within the study area.
Camera method Reducing survey effort without compromising detection proba-
bilities has many obvious advantages (MacKenzie and Royle
Fig. 2. Komodo dragon (a) detection probability and (b) site occupancy
2005). Most importantly, saved survey effort can be allocated
estimated from baited or unbaited cameras deployed for 6- or 30-day
sampling events respectively. The bars report the mean estimate, and the
into additional sites, survey visits or additional study areas in
upper and lower 95% confidence intervals are indicated by the error bars. different ways (Sewell et al. 2012). From our perspective, the
biggest advantage is that reduced survey effort can be invested
into additional camera monitoring activities for more broadly
1.0 assessing the conservation status of Komodo dragon popula-
Cumulative detection probaility

tions. For example, we have recently used baited camera

0.8
0.6
0.4
0.2
0.0
0 1000
Sampling effort (days)
Fig. 3. The cumulative detectability curves for Komodo dragons estimated
from baited and unbaited camera occupancy models. These curves represent
the probability that Komodo dragons will be detected at least once with each
treatment after sequential 1-week sampling period at each camera trap where
Komodo dragons are present. The lines report the mean estimate and the
dashed upper and lower lower lines are the 95% standard error of the mean.
monitoring surveys beyond this study area to evaluate the
distribution of the Komodo dragon across Flores (,400 moni-
toring stations across 1200 km of coastline; Ariefiandy et al.
2021). This feat would not have been possible without using
baited cameras to achieve high Komodo dragon detection
relative to their survey effort requirements.
It is argued that the use of attractants to increase a species
Baited camera
Unbaited camera
detection must be considerate of any effects on monitoring
estimates and arising inference (du Preez et al. 2014). For
example, if increased estimates of detection at baited cameras
2000 3000 4000
arise because of bait effects on animal space-use or daily
movements, it could bias parameter estimates. In the case of
baited cameras, baits could increase residency times or attract
animals beyond their normal home-range area to inflate esti-
mates of detection probability and site occupancy (Stewart et al.
2019). This problem could be especially acute if individual
animals, particularly those in low-density populations, are
detected at multiple camera stations beyond their home range.
300 Wildlife Research
Consequently, ensuring spatial independence for camera data is
a crucial aspect of monitoring design (Meek et al. 2014;
O’Connell et al. 2010; Geyle et al. 2020). We know that the
independence of data among camera monitoring sites is largely
met for Komodo dragon, because our prior mark–recapture-
based studies using traps with similar inter-site distances
resulted in a ,10% within-study recapture rate of individuals
(Ariefiandy et al. 2013, 2014).
This study also indicated that estimates of Komodo dragon
site occupancy recorded within the Wae Wuul Nature Reserve
are significantly lower than those generally recorded for popula-
tions in the adjacent Komodo National Park (Purwandana et al.
2014b; Ariefiandy et al. 2015). Adult Komodo dragons, as apex
predators, mainly prey on ungulates, particularly Rusa deer
(Rusa timorensis), wild pig (Sus scrofa), and, in some locations,
water buffalo (Bubalus bubalis; Auffenberg 1981; Bull et al.
2010; Purwandana et al. 2016). Thus, we attribute this lower
occupancy estimate to be in part associated with the commen-
surate reduction of large ungulate prey availability on Flores
(Ariefiandy et al. 2011, 2015, 2016; Jessop et al. 2020). Reduced
prey is a presumed consequence of historical and increasingly
contemporary human-mediated processes (e.g. fire, poaching,
invasive predators) affecting Komodo dragon habitats on Flores
(Ariefiandy et al. 2020).
Here we advocate that protected-area enhancement actions
and community conservation approaches are needed to address
the current threats to Komodo dragons on Flores. For example,
unlike Komodo National Park, the Wae Wuul Nature reserve is
comparatively under-resourced in staff and logistical resources.
Thus, aside from ongoing monitoring of Komodo dragon popu-
lations, it is necessary to ensure that integrative conservation
actions are used to ensure prey and predator persistence in this
reserve (Ariefiandy et al. 2015, 2020). Thus, this reserve could
benefit from additional infrastructure (e.g. ranger posts) and
increased patrolling and surveillance measures that would
benefit both Komodo dragons and their ungulate prey (Hilborn
et al. 2006; Ariefiandy et al. 2015, 2020). However, as human
activities increasingly modify the habitats that directly border
this reserve, community-based conservation actions must also
be implemented in neighbouring communities (Ariefiandy et al.
2015, 2020). For example, implementing conservation aware-
ness meetings in local communities to inform and discuss the
value of protecting natural values within this reserve are deemed
essential (Kamil et al. 2019). Furthermore, working with com-
munities to reduce rates of incursions of village dogs or livestock
and stopping villagers from setting fire to habitats within or
adjacent to the reserve could be important steps to promote the
conservation of Komodo dragons in this key protected area on
Flores (Ariefiandy et al. 2020).
In conclusion, our study demonstrated that optimising cam-
era survey methods using baits compared with unbaited cameras
can provide a better method for estimating Komodo dragon
occupancy. This result was particularly important in this study
because we aimed to effectively monitor a very low-density
population in a key protected area on Flores. We believe our
baited camera monitoring approach now lends itself to under-
standing population responses to ecological and anthropogenic
factors, hence informing conservation efforts in this nature
reserve (Ariefiandy et al. 2015).
D. Purwandana et al.
Conflicts of interest
The authors declare that they have no conflicts of interest.
Author contributions
Study design and fieldwork: DP, AA, MA, SAN, MS and TSJ;
data analysis: TSJ; writing: TSJ.
Acknowledgements
We are grateful to the Directorate General of Conservation of Natural
Resources and Ecosystem (DITJEN KSDAE) and Eastern Lesser Sunda
Central Bureau for Conservation of Natural Resources (BBKSDA NTT) for
issuing research permits and the BBKSDA NTT authority and members of
staff and local community volunteers for support and their enthusiastic
involvement in the project. Funding for fieldwork (2014–2019) was pro-
vided through the European Association of Zoos and Aquaria, Chester Zoo,
and the Ocean Park Conservation Foundation Hong Kong.
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